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Applications of Genetics to Arthropods of Biological Control Significance Edited by S. Karl Narang Alan C. Bartlett Robert M. Faust Boca Raton London New York CRC Press CRC Press is an imprint of the Boca RaTatyolonr & FAranncnis AGrorubpo, arn inLfoormnad bounsin essTokyo First published 1993 by CRC Press Taylor & Francis Group 6000 Broken Sound Parkway NW, Suite 300 Boca Raton, FL 33487-2742 Reissued 2018 by CRC Press © 1993 by CRC Press, Inc. CRC Press is an imprint of Taylor & Francis Group, an Informa business No claim to original U.S. Government works This book contains information obtained from authentic and highly regarded sources. Reasonable efforts have been made to publish reliable data and information, but the author and publisher cannot assume responsibility for the validity of all materials or the consequences of their use. The authors and publishers have attempted to trace the copyright holders of all material reproduced in this publication and apologize to copyright holders if permission to publish in this form has not been obtained. If any copyright material has not been acknowledged please write and let us know so we may rectify in any future reprint. Except as permitted under U.S. Copyright Law, no part of this book may be reprinted, reproduced, transmitted, or utilized in any form by any electronic, mechanical, or other means, now known or hereafter invented, including pho- tocopying, microfilming, and recording, or in any information storage or retrieval system, without written permission from the publishers. For permission to photocopy or use material electronically from this work, please access www.copyright.com (http:// www.copyright.com/) or contact the Copyright Clearance Center, Inc. (CCC), 222 Rosewood Drive, Danvers, MA 01923, 978-750-8400. CCC is a not-for-profit organization that provides licenses and registration for a variety of users. For or- ganizations that have been granted a photocopy license by the CCC, a separate system of payment has been arranged. Trademark Notice: Product or corporate names may be trademarks or registered trademarks, and are used only for iden- tification and explanation without intent to infringe. Library of Congress Cataloging-in-Publication Data Applications of genetics to arthropods of biological control significance / editors, S. Karl Narang, Alan C. Bartlett, Robert M. Faust. p. cm. Papers presented at a workshop held at the 19th International Congress of Entomology. Beijing, China. 1992. Includes bibliographical references and index. ISBN 0-8493-2607-9 1. Arthropods as biological pest control agents—Congresses. 2. Arthropods—Genetics— Congresses. I. Narang, S. Karl. II. Bartlett, Alan. C. Ill. Faust, Robert, M. IV. International Congress of Entomology (19th: Peking, China: 1992) SB976.A78A66 1994 632’.7—dc20 93-44973 A Library of Congress record exists under LC control number: 93044973 Publisher’s Note The publisher has gone to great lengths to ensure the quality of this reprint but points out that some imperfections in the original copies may be apparent. Disclaimer The publisher has made every effort to trace copyright holders and welcomes correspondence from those they have been unable to contact. ISBN 13: 978-1-315-89066-1 (hbk) ISBN 13: 978-1-351-06976-2 (ebk) Visit the Taylor & Francis Web site at http://www.taylorandfrancis.com and the CRC Press Web site at http://www.crcpress.com Big~cycd bug, Geocoris punctipes (Say). Photo courtesy of Jack Dykinga. EDITORS S. Karl Narang, Ph.D. is a Research Leader of the Insect Genetics and Cellular Biology Research Unit at the U.S. Department of Agriculture, Agricultural Research Service, Fargo, ND. Dr. Narang received his genetics training at the University of Illinois, Urbana-Champaign, receiving the M.S. degree in 1968 and the Ph.D. degree in 1970. He served as a Senior Scientist/Geneticist at the World Health Organization's Project on Genetic Control (New Delhi) in 1971; as Associate Professor and later Chairman of the Department of Animal Biology, University of Brasilia, Brasilia, Brazil from 1972 to 1979; Research Scientist at the Department of Entomology, University of Florida, Gainesville from 1980 to 1985; and Research Geneticist at the USDA, ARS laboratory in Gainesville from 1986 to 1989. It was in 1990 that he assumed his present position. Dr. Narang is a member of the Genetic Society of America, American Genetics Society (life member), Entomological Society of America, Florida Entomological Society (serving as subject-editor in genetics), American Mosquito Control Associa- tion, Panj ab University Zoological Society (life member), has been a member of the American Association of Advancement of Science, New York Academy of Science, Sigma Xi, Brazilian Genetics Society, Brazilian Society for the Advancement of Science, and Brazilian Biomedical Association. He has served as a consultant and technical advisor to the National Academy of Science, National Research Council, Board of Science and Technology for International Development, and reviewed overseas projects on genetics and ecology of insect pests. He has served as consultant to the U.S.-Brazil Biomedical Research Collaboration, University of Maryland and University of Bahia, Salvador, Bahia, Brazil. He has received several awards includ- ing Gold medals for highest academic achievements during his B.S. and M.Sc. studies, Fullbright Travel award, World Health Organization Fellowship award, and WHO's WHO in Frontiers of Science and Technology (1985, 2nd edition of Marquis WHO's WHO). He has been the recipient of many research grants for the National Research Council and Walter Reed Army Institute of Research. He has organized many workshops and chaired symposia sponsored by FAO/IAEA/DOE/USDA. the Rockefeller Foundation, and by many national and international societies. Dr. Narang is the author of more than 120 papers and has co-edited 2 books and contributed 8 book chapters. His current major research interests relate to population genetic structures of insect pests and genetic engineering of insects as they relate to implementation of biological control programs. Alan C. Bartlett, Ph.D., is a Research Geneticist (insects) at the U.S. Department of Agriculture, Agricultural Research Service, Western Cotton Research Laboratory, Phoenix, AZ. Dr. Bartlett received an Associate in Science degree from Carbon College (now College of Eastern Utah), Price, UT (1954), his B.A. (cum laude) and M.S. degrees in genetics at the University of Utah, Salt Lake City, UT (1956, 1957) and his Ph.D. in population and radiation genetics at Purdue University, Lafayette, IN (1962). He was an instructor at Carbon College in 1957-1958 and at Purdue University in 1960--1962. He joined USDA/A RS as an insect geneticist in 1962 at the Boll Weevil Research Laboratory, State College (now Mississippi State University), MS. In 1967 he transferred to the Cotton Insects Research Laboratory in Tucson, AZ and in 1970 he assumed his present position as Research Geneticist in Phoenix, AZ. He served as an adjunct professor of genetics at Mississippi State University and Arizona State University, Tempe, AZ. Dr. Bartlett is a member of the Genetics Society of America, Entomological Society of America, Southwestern Entomological Society, Sigma Xi, and American Institute of Biological Sciences, and has been a member of AAAS, American Genetic Association, Phi Sigma, and the Arizona Academy of Science. He is listed in Marquis Who's Who in America, Who's Who in Science and Technology, and Who's Who in the West. He has organized and participated in symposia sponsored by the Entomo~ logical Society of America, International Atomic Energy Agency, Hungarian Acad- emy of Science, International Crops Research Institute for the Semi-Arid Tropics, USDA, ARS, the Arizona Academy of Sciences, and the International Congress of Entomology . He served as president of the Mississippi State University Chapter of Sigma Xi and chairman of the Tempe Gifted Association. He received the Outstand- ing Alumnus award from the College of Eastern Utah in February 1993. Dr. Bartlett is the author of more than 80 papers, the co-editor of 1 book, and has contributedchapters to 8 books. His current major research interests are the develop- ment of genetic methods of control of insects that attack cotton, the development of genetic methods of identification of geographic or host races of insects, and the genetic effects of the domestication process on insects. Robert M. Faust, Ph.D., is a National Program Leader of fundamental and molecu- lar biology on the National Program Staff of the U.S. Department of Agriculture, Agricultural Research Service, Beltsville, MD. Dr. Faust received his training in microbiology and entomology at the University of Maryland, College Park, receiving his B.S. in 1963, his M.S. in 1966, and Ph.D. in 1970. He served at the USDNARS, Market Quality Research Laboratory in 1959-1960, the Bioenvironmental Bee Labo- ratory, 1960--1988, and has been in his present position since 1988. Dr. Faust is a founding member of the Society of Invertebrate Pathology, a member of the American Society of Microbiology, the Entomological Society of America, AAAS, New York Academy of Science, Japanese Society of Sericultural Science, Biochemical Society of the Republic of Korea, Korean Society of Applied Microbiology, American Registry of Professional Entomologists, American Legion, and Sigma Xi. Dr. Faust was a visiting scientist at the Armenian Academy of Science Institute of Microbiology, USSR, 1976; Korean Institute of Science and Technology, Seoul, 1985; part-time faculty of Prince Georges Community College, Largo, MD, 1964--1984; and president, SCH Corporation, 1977. He served on the executive board of the Center of Biological Control of Mosquitoes, Ben Gurion University, 1981. Dr. Faust has served on the editorial board of the Journal of Invertebrate Pathology ( 1980--1985) and authored books and articles on such varied subjects as Diseases of Insects, 1974; Microbial and Viral Insecticides, 1982; Genetic Engineering in the Plant Sciences, 1981; The Molecular Biology of the Bacilli, 1985; and contributed critical reviews and scientific articles to professional journals. PREFACE The continued use of broad-spectrum synthetic chemical insecticides over more than four decades has led to widespread insect resistance, as well as heightened the concern over their adverse effect on the environment and human health. Further, the U.S. Environmental Protection Agency has banned the use of many such chemicals and severely restricted the use of others; more are likely to be banned. Consequently, the discovery and development of environmentally sound, publicly acceptable, and viable economical strategies of pest control have taken on a new urgency. An important foundation of pest control is integrated pest management (IPM), of which an important component includes the use of biological control agents; IPM offers one of the best long-term strategies to help reduce or eliminate the use of synthetic chemical pesticides. Biological control is based on the use of selected natural enemies from the native range of the pest to reduce the pest's negative effects. Generally, it includes (1) importation and release of natural enemies, (2) augmentation of populations of established biotypes by laboratory mass rearing, and (3) conservation of natural enemies by manipulation of the habitats. However, biological control technology is considered by many to be still in its infancy. The literature contains many examples of both successes and failures. The failure of biological programs to reduce the intended effect can be traced to an inadequate knowledge and understanding of fundamental pest life processes, biosystematics, genetics, ecological influences; and pest, natural enemies, host interrelationships. Thus, there are many questions that must be resolved before biological control strategies can reach a higher level of success as an integral component of IPM-based systems approaches to pest control. First, it is critical that we know the specific status of each target pest and the natural enemies of that pest. For example, many parasitic species have not yet been identified and named, and many of these are sibling species. The presence of unrecognized species in a collection might account for the partial (or even lack of) success of some biocontrol programs. Some investigators question the need for concern about genome diversity in biocontrol organisms. The identification of cryp- tic species is important in two respects. Different species may attack different hosts. If they do attack the same host, they may differ in their attributes for biological control. There are many examples where failure to detect cryptic taxa in the target pest or its biocontrol agents has led to failure of biocontrol projects. It should be evident to the reader that detecting cryptic taxa is not a trivial matter, it can mean the difference between success and failure. Populations of the same species may differ in relevant biological characteristics important to biological control. Therefore, biological and genetic variation within and among species must be characterized to develop effective biocontrol strategies. Such studies become even more critical when new exotic enemies are imported and their establishment in the new environment forms the basis of the program. Many areas and crops have been or are currently being treated with pesticides that can destroy the parasitoid as effectively as it does the pest species. One way to solve this problem would be to use parasitoids that are resistant to the pesticide. This would allow the release of parasitoids in combination with pesticide application, thus increasing the effectiveness of both. Extensive work is being done in a few labora- tories to select for resistant strains from field populations. This is an expensive and laborious task and has been criticized for this reason. Development of pesticide- resistant strains is one avenue that may be exploited using biotechnological tech- niques. Augmentation of natural enemies involves mass production and periodic coloni- zation under laboratory conditions. When a colony is established from a sample of a natural population collected abroad, only some of the genetic variability of the area of origin is represented. This may account for the partial success of some biocontrol agents, but some investigators question the need to be concerned about genetic diversity. Many, if not most, of the highly successful biological control projects have occurred from limited introduction that would appear to represent only a fraction of species genome diversity. This would suggest that other factors, such as exclusion of pathogens, might take priority over concern about genome diversity as exotic natural enemies are chosen and prepared for field release. The successful establishment and spread of the vedalia beetle, Rodolia cardinalis, in California from one introduction is the example often cited in support of this view. That success has been repeated often following introduction of the vedalia beetle from California into other coun- tries. Often, a small subset of the original California introduction was included in the subsequent releases. Contrary to the success of the vedalia beetle in some parts of California is the failure or only partial success of this predator in coastal California and Bermuda. Was genome diversity too limited for the insect to perform adequately under cooler climatic conditions? Are there biotypes which would perform better under cooler climates? There are no easy answers to such questions. Adequate data is lacking to assess the breadth of genomic diversity necessary to ensure effectiveness of natural enemies. Therefore, development of methods to determine the role of genome diversity (and to increase it, if necessary) in the effectiveness of natural enemies should be done to ensure success of biological control strategies. Recent advances in biotechnology provide tremendous potential for improvement of the effectiveness of natural enemies against insect pests. This could range from the development of genetic sexing strains to engineering of disease and pesticide resis- tance for beneficial insects. However, genes controlling desirable traits must be identified. Next, these genes must be cloned and characterized. Finally, these genes must be suitably modified and successfully introduced into the target organism's DNA such that the desired trait will be transmitted in a Mendelian fashion. It was against this background that a symposium/workshop on "Genetic Applica- tions to Arthropods of Biological Control Significance" was held at the XIX Inter- national Congress of Entomology in Beijing, China in the summer of 1992. This symposium/workshop brought together scientists from federal and state research institutions, both foreign and domestic, to review the state of knowledge, to identify research gaps and needs, and to suggest corresponding approaches to meet the needs of this important approach to the use of parasitoids and predators in biological control. It was recognized that there is a paucity of information on the genetics of natural enemies in the literature. Therefore, this book compiles papers presented during the symposium/workshop and also articles by other contributors who were unable to attend. We believe this book details the current state of knowledge and makes an earnest attempt at addressing many of the basic issues in terms of genetic innovations that have the potential to considerably improve the effectiveness of modern biological control programs. S. K. Narang A. C. Bartlett R. M. Fanst

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